异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

Heterostyly accelerates diversification via reduced extinction in primroses

The exceptional species diversity of flowering plants, exceeding that of their sister group more than 250-fold, is especially evident in floral innovations, interactions with pollinators and sexual systems. Multiple theories, emphasizing flower–pollinator interactions, genetic effects of mating systems or high evolvability, predict that floral evolution profoundly affects angiosperm diversification. However, consequences for speciation and extinction dynamics remain poorly understood. Here, we investigate trajectories of species diversification focusing on heterostyly, a remarkable floral syndrome where outcrossing is enforced via cross-compatible floral morphs differing in placement of their respective sexual organs. Heterostyly evolved at least 20 times independently in angiosperms. Using Darwin''s model for heterostyly, the primrose family, we show that heterostyly accelerates species diversification via decreasing extinction rates rather than increasing speciation rates, probably owing to avoidance of the negative genetic effects of selfing. However, impact of heterostyly appears to differ over short and long evolutionary time-scales: the accelerating effect of heterostyly on lineage diversification is manifest only over long evolutionary time-scales, whereas recent losses of heterostyly may prompt ephemeral bursts of speciation. Our results suggest that temporal or clade-specific conditions may ultimately determine the net effects of specific traits on patterns of species diversification.     

The different forms of flowers – what have we learned since Darwin?

Darwin's book, The Different Forms of Flowers on Plants of the Same Species, has stimulated an extraordinary amount of original research since its publication in 1877. In his book, Darwin focused primarily on heterostylous reproductive systems in flowering plants, in which two or three reproductive morphs with reciprocal placement of anthers and stigmas occur in populations. These morphs are usually self‐incompatible and cross‐incompatible with individuals possessing the same reproductive morph. Many of the papers on heterostyly published since Forms of Flowers appeared have focused on the questions raised by Darwin about the evolution and function of heterostyly. Darwin's hypothesis that heterostyly promotes cross‐pollination between different morphs has been largely substantiated, despite the difficulties in finding the ideal experimental system to address this question. Heterostyly is now known to occur in many more plant families than at the time Forms of Flowers was published and, as expected, the heterostylous syndrome is now defined more broadly than in Darwin's time. The origin of heterostyly remains an area of active research, with hypotheses stressing either the evolution of heteromorphic self‐incompatibility as the first step in the evolution of this reproductive system or, alternatively, the evolution of the reciprocal features of floral morphology. Phylogenetic approaches, combined with studies on the physiological and molecular genetic basis of heterostyly, offer promise in helping to resolve questions about the origin of heterostyly. There is no doubt that heterostyly has evolved on multiple occasions and that self‐incompatibility associated with heterostyly is unrelated to the more common multi‐allelic self‐incompatibility systems found in monomorphic species. Further progress in understanding conditions favouring evolution of heterostyly will depend on an increased understanding of the relation between the reciprocal morphological features of the breeding system and the nature of self‐incompatibility. Almost a century and a half after the appearance of The Different Forms of Flowers on Plants of the Same Species, heterostyly remains an active area of research. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 249–261.     

Distyly and heteromorphic incompatibility in oceanic island species ofErythroxylum (Erythroxylaceae)

Surveys of oceanic island floras have shown that heterostyly is usually absent in such regions, probably because this floral polymorphism is often associated with a self-incompatibility system. In this context we describe the floral biology of three species ofErythroxylum on La Réunion island and examine the compatibility relationships of one of these species,E. laurifolium. All three species are distylous but differ in relative stigma-anther separation in the different morphs. In general, short-styled flowers have greater stigma-anther separation than long-styled flowers, which are often homostylous in appearance. This lack of stigma-anther separation in long-styled flowers is due to style twisting which improves reciprocity at the high organ level. The reduced stigma-anther separation does not appear to be associated with the evolution of selfing asErythroxylum laurifolium shows heteromorphic self-incompatibility. The presence of heteromorphic incompatibility in a group of species that have colonized an oceanic island is discussed.     

Importance of Queen Bumble Bees as Pollinators Facilitating Inter-morph Crossing in Primula sieboldii

Abstract Flower-visiting insects, the distribution of pollen on an insect's body, and fruit and seed sets of Primula sieboldiiE. Morren were investigated in a maritime deciduous forest habitat of the Hidaka region in Hokkaido. Queens of Bombus diversus tersatus Smith (Hymenoptera: Apidae), Bombus schrencki albidopleuralis Skorikov, and Bombus deuteronymus deuteronymus Schulz were observed to visit the flowers of the species. Bumblebee claw marks were found on the flower petals of 68% of the flowering ramets of the species. Scanning-electronmicroscopy revealed that pin and thrum pollens of P. sieboldii were deposited in different positions along the proboscis of a B. diversus tersatus queen (n=38400 grains). On the glossa, all the observed pollen was of the pin type, while more than half of the pollen grains observed on the upper part of the proboscis, galea, stipes, and maxillary palpus were of the thrum type. In the field, high fruit and seed sets were recorded for both pin and thrum morphs, indicating the occurrence of sufficient legitimate pollination during the flowering season. In addition to the effectiveness of queen bumblebees as pollinators for P. sieboldii suggested in this study, phenological and morphological matches, i.e., early spring flowering during the queen bee emergence season and the similar lengths of the corolla tube and the bee proboscis, also support the inference that queen bumble bees play an important role in legitimate pollination of P. sieboldii.